Etruscan rhinoceros
| Etruscan rhinoceros | ||||||||||||
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Etruscan rhinoceros skull |
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| Temporal occurrence | ||||||||||||
| Old to earliest Middle Pleistocene | ||||||||||||
| 2.588 million years to 700,000 years | ||||||||||||
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| Systematics | ||||||||||||
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| Scientific name | ||||||||||||
| Stephanorhinus etruscus | ||||||||||||
| ( Falconer , 1868) | ||||||||||||
The Etruscan rhinoceros ( Stephanorhinus etruscus ) is an extinct rhinoceros species that lived in Europe mainly in the Old Pleistocene and the earliest Middle Pleistocene 2.5 to 0.7 million years ago. It belongs to the group of two-horned rhinos of Eurasia, which is closely related to the Sumatran rhinoceros ( Dicerorhinus sumatrensis ) that lives in Southeast Asia today but is critically endangered . The vast majority of the Etruscan rhinoceros habitat were open forest landscapes, where it fed on soft plant food.
distribution and habitat
The Etruscan rhinoceros preferred largely sub-Mediterranean to temperate climates . Its main distribution area was western and southern Europe , where it occurred in the area of what is now France , on the Iberian , Apennine , Balkan Peninsula and the Peloponnese . To the east its distribution area extended to the Caucasus . The areas north of the Alps may only have been reached during the warmer phases of the Old Pleistocene (around 2.5 to 0.8 million years ago); it is far less proven here. The northernmost occurrences are known so far near the North Sea in today's Netherlands (Tegelen) and on the British Isles (East Runton).
In western and southern Europe, the Etruscan rhinoceros initially coexisted with the great rhino representatives Stephanorhinus jeanvireti and Stephanorhinus megarhinus . Especially in Eastern Europe, in the Chapri fauna complex of the Old Pleistocene, it is associated with early representatives of Elasmotherium , an elephant-sized rhinoceros. In its later phase, the rhinoceros species shared their habitat with the Hundsheimer rhinoceros ( Stephanorhinus hundsheimensis ). Numerous fossils of North Alpine, Central European sites from the transition from the Old to the Middle Pleistocene, which were originally associated with the Etruscan rhinoceros, belong to this more modern rhino form. Other frequently associated fauna elements are the southern elephant and bison schoetensacki .
Physique and lifestyle
The Etruscan rhinoceros was a medium-sized representative of the rhinos and reached a head-trunk length of 250 cm with a shoulder height of around 160 cm. The weight is put at between 400 and 750 kg, with which the rhino species reached about the size of today's Sumatran rhinoceros ( Dicerorhinus sumatrensis ).
The skull of the Etruscan rhinoceros was about 63 cm long. The occiput was not pulled out much and was shaped almost at right angles. This led to a relatively horizontal head posture, just as in the fossil forest rhinoceros ( Stephanorhinus kirchbergensis ) and in today's Sumatran rhinoceros. It had a weak bulge as a base for the neck muscles. The nasal bone was rather poorly developed and only ossified in the anterior area, but had a rough, pearl-like surface that indicates the position of the anterior horn. The smaller, second horn sat on the forehead and was indicated by similar surface structures. The interior of the nose reached to the posterior premolar . The eye socket was above the third molar .
The lower jaw was built gracefully and was 49 cm long. He had a long but narrow symphysis . As with the other Stephanorhinus representatives, the anterior dentition was reduced, while the posterior had only receded the anterior premolar and consisted of three premolars and three molars per jaw branch. The dental formula was accordingly: . In individual individuals, however, the first premolar can still be developed. The molars were clearly low-crowned ( brachyodont ). The premolars and front molars had a more square to rectangular outline, the rearmost molar was roughly triangular in shape. The second molar represented the largest tooth in the dentition.
The torso skeleton is quite well known due to numerous finds. The spine comprised - based on a largely complete skeleton found near the central Italian town of Terni - Stephanorhinus - typically at least 7 cervical, 18 thoracic and 4 lumbar vertebrae. The limbs were very slender, even slimmer than the roughly equal but heavier Hundsheimer rhinoceros ( Stephanorhinus hundsheimensis ). The humerus reached 39 cm in length, the ulna 48 cm. The thigh bone was 44 cm long , the shin 37 cm long. The three-pronged hands and feet had a massive central ray (Metapodium III), whereby the metacarpal bone was 20 cm long and the metatarsal bone was 18 cm long.
The horizontal head posture and the low-crowned teeth suggest that the Etruscan rhinoceros largely nourished itself on soft vegetable foods ( browsing ). As a result, leaves , bark , flowers or buds should have been the main sources of food. However, it cannot be ruled out that a small proportion of hard grass food was also consumed. However, comparable to today's Indian rhinoceros ( Rhinoceros unicornis ), it should have accounted for far less than 50% of all plants consumed. It is possible that the rhino species lived in semi-open forest landscapes, which is also indicated by the relatively slender limbs.
Systematics and research history
The Etruscan rhinoceros belongs to the genus Stephanorhinus , a group of largely Eurasian representatives of rhinoceros that occurred from the Miocene to the Pleistocene and whose closest living relative is the Sumatran rhinoceros . The relationships within the Stephanorhinus line are still largely unknown, two lines are generally assumed: Stephanorhinus megarhinus - Stephanorhinus kirchbergensis and Stephanorhinus etruscus - Stephanorhinus hundsheimensis - Stephanorhinus hemitoechus . However, the closer connection between Etruscan and Hundsheimer rhinoceros is also partly doubted, as the latter has much more archaic molars. The ancestor of the Etruscan rhinoceros was Stephanorhinus jeanvireti . Both may represent immigrants from Asia in Europe.
Most researchers describe the Etruscan rhinoceros as relatively unchangeable in its phylogenetic development. However, as early as 1934, Friedrich Zeuner pointed to the increase in size of the anterior horn base in relation to the length of the skull, which is particularly noticeable in the latest representatives. By Claude Guérin then two subspecies, were S. e. etruscus and S. e. brachycephalus introduced. The later form S. e. brachycephalus , which largely originates from the middle Pleistocene , is now regarded as a Hundsheimer rhinoceros, which , according to anatomical studies, actually had a larger nasal horn. According to Paul Mazza , the Etruscan rhinoceros is identical to the Chinese variant Stephanorhinus yunchuchenensis , which would extend the range of this species to East Asia. So far, the individual members of the genus Stephanorhinus , especially the East Asian, were subject to no major revision.
The first description of the Etruscan rhinoceros as Rhinoceros etruscus was written by Hugh Falconer in 1868 , but his report was published posthumously . However, Falconer had already mentioned the name in a letter to David Thomas Ansted in 1859 . The description was based on fossils from Tuscany , from which the species name as a former settlement area of the Etruscans is derived. Due to the later recognized, closer relationship to the Sumatran rhinoceros - unifying features are the two developed horns and the partly ossified nasal bone - the genus name was changed to Dicerorhinus . In 1942, Miklós Kretzoi introduced the genus name Stephanorhinus , which is valid today and which he established due to the differences in the front teeth between the Sumatran rhinoceros and the Pleistocene rhinos of Eurasia.
Tribal history
The Etruscan rhinoceros first appeared in the early Pleistocene, possibly even in the late Pliocene . In Europe it appears almost simultaneously with the equine Equus stenonis . One of the oldest sites is that of Chilhac in France , which is assigned to the Gelasium . At that time, the rhinoceros species was common in western Europe, but it was also found in eastern Europe. The rhinoceros species first appeared in Central Europe in the late Old Pleistocene and can be found in the Erpfinger Cave in Baden-Württemberg , among other places . The last appearance of the Etruscan rhinoceros is in Atapuerca in Spain , a site that is also associated with important early human fossils (see also Homo antecessor ) and legacies, more than 700,000 years ago.
Individual evidence
- ↑ a b c Ralf-Dietrich Kahlke, Nuria García, Dimitris S. Kostopoulos, Frédéric Lacombat, Adrian M. Lister, Paul PA Mazza, Nikolai Spassov, Vadim V. Titov: Western Palaearctic palaeoenvironmental conditions during the Early and early Middle Pleistocene inferred from large mammal communities, and implications for hominin dispersal in Europe. In: Quaternary Science Review. 2010, pp. 1–28.
- ↑ Walter Steiner: Europe in the primeval times. Munich 1993, ISBN 3-576-10276-0 .
- ^ Jean-Philip Brugal, Roman Croitor: Evolution, ecology and biochronology of herbivore associations in Europe during the last 3 million years. In: Quaternaire. 18 (2), 2007, pp. 129-152.
- ↑ a b Friedrich E. Zeuner: The relationships between skull shape and way of life in recent and fossil rhinos. In: Reports of the Natural Research Society in Freiburg. 34, 1934, pp. 21-80.
- ↑ a b Mikael Fortelius, Paul Mazza, Benedetto Sala: Stephanorhinus (Mammalia: Rhinocerotidae) of the Western European Pleistocene, with a revision of S. etruscus (Falconer, 1868). In: Palaeontographia Italica. 80, 1993, pp. 63-155.
- ↑ a b c H. Loose: Pleistocene Rhinocerotidae of W. Europe with reference to the recent two-horned species of Africa and SE Asia. In: Scripta Geolica. 33, 1975, pp. 1-59.
- ↑ Oierluigi Ambrosetti: Lo scheletro di Dicerorhinus etruscus (Falc.) Di Capitore (Umbria meridional). In: Geologica Romana. Roma 11, 1972, pp. 177-198.
- ↑ a b c Paul Mazza: The Tuscan Early Pleistocene rhinoceros Dicerorhinus etruscus. In: Palaeontographia Italica. Pisa 75, 1988, pp. 1-87.
- ^ Paul Palmqvist, Darren R. Grocke, Alfonso Arribas, Richard A. Fariña: Paleoecological reconstruction of a lower Pleistocene large mammal community using biogeochemical (d13C, d15N, d18O, Sr: Zn) and ecomorphological approaches. In: Paleobiology. 29 (2), 2003, pp. 205-229.
- ↑ a b Frederic Lacombat: Phylogeny of the genus Stephanorhinus in the Pliocene-Pleistocene of Europe. In: Hallesches Jahrbuch für Geoswissenschaften. 23, 2007, pp. 63-65.
- ↑ a b c d Jan van der Made: The rhinos from the Middle Pleistocene of Neumark-Nord (Saxony-Anhalt). In: Dietrich Mania et al. (Ed.): Neumark-Nord: An interglacial ecosystem of the Middle Paleolithic man. Halle / Saale 2010, ISBN 978-3-939414-37-7 , pp. 433-527. (Publications of the State Museum for Prehistory 62)
- ↑ Claude Guérin: Les Rhinocerotidae (Mammalia, Perissodactyla) du Miocène terminal au Pléistocène supérieur d'Europe occidentale comparés aux espèces actuelles: tendances évolutives et relations phylogénétiques. In: Géobios. 15, 1982, pp. 599-605.
- ↑ Odile Boeuf: Le Dicerorhinus etruscus (Rhinocerotidae, Mammalia) du site Pliocène supérieur de Chilhac (Haute-Loire, France). In: Géobios. 28 (3), 1995, pp. 383-391.